Three new goatfishes of the genus Upeneus (Mullidae) from the Indo-Pacific, with a redescription of colour patterns in U. margarethae Author Uiblein, Franz Vietnam National Museum of Nature, Vietnam Academy of Science and Technology (VAST), Ha Noi, Vietnam franz@hi.no Author Gledhill, Daniel C. Division of Water and Marine Resources, Department of Primary Industries, Parks, Water and Environment, Hobart, Australia Biological Faculty, Moscow State University, Moscow, Russia Author Pavlov, Dimitri A. Author Hoang, Tuan Anh Graduate University of Science and Technology, VAST, Ha Noi, Vietnam Author Shaheen, Shaker Faculty of Marine Sciences and Fisheries Red Sea University, Port Sudan, Sudan text Zootaxa 2019 2019-10-08 4683 2 151 196 journal article 22524 10.11646/zootaxa.4683.2.1 30220eba-fa3d-4311-84b0-07b0d8ba23d0 1175-5326 3772664 4365B195-E427-421E-9C4D-6DB746F66B8A Upeneus gubal n. sp. Uiblein Gubal goatfish ( Figures 1 , 5 , 7 ; Tables 1 , 5 , 6 ) Holotype . MNHN 2011-1515 , adult, 86 mm SL, WIO , Red Sea , Gulf of Suez , Strait of Gubal , N of Gubal Island , 27° 49’ 59” N , 33° 43’ 1” E , 70 m depth , 28 Dec. 1928 , collector Robert P. Dollfus. Diagnosis. Dorsal fins VIII + 9, the first spine minute; pectoral fins 15; gill rakers 6 + 19 = 25; lateral-line scales 28–29; measurements in % SL: body depth at first dorsal-fin origin 24; body depth at anus 21; caudal-peduncle depth 11; caudal-peduncle width 3.3; maximum head depth 20; head depth through eye 16; suborbital depth 9.0; interorbital length 8.2; head length 29; snout length 11; postorbital length 11; orbit length 7.4; upper-jaw length 11; barbel length 19; interdorsal distance 16; caudal-peduncle length 24; caudal-fin length 31; anal-fin height 19; pelvic-fin length 22; pectoral-fin length 22; pectoral-fin width 4.8; first dorsal-fin height 22; second dorsal-fin height 19; preserved colour pale brown. Description. Measurements in % SL and counts are given in Table 1 ; morphometric data as ratios of SL for holotype : body elongate, body depth at first dorsal-fin origin 4.2, body depth at anal-fin origin 4.8, head length 3.4, larger than maximum body depth and subequal to caudal-fin length (3.2); second dorsal-fin height 5.2, shallower than first dorsal fin (4.4) and similar to barbel length (5.4); pelvic-fin length 4.5, equal to length of pectoral fins (4.5) and similar to body depth at first dorsal-fin origin; caudal-peduncle depth 9.2, much larger than orbit length (14); and caudal-peduncle width 30, much narrower than pectoral-fin width (21). Colour . Preserved fish . ( Figure 7 D ). Body and head entirely pale brown, with brown patches or blotches on head below and behind eyes and on anterior ventral side of body; barbels pale creamy; no remains of dark pigmentation deriving from caudal-fin bars, dark mid-lateral dots, saddle behind second dorsal fin, or second dorsal-fin distal stripe; fins pale brown, partly hyaline. FIGURE 8. Three selected morphological characters against SL in Upeneus caudofasciatus n. sp. The 65 mm demarcation for subadults and adults is indicated by a dotted black dash. TABLE 8. Morphometric, meristic and preserved colour characters in types of Upeneus caudofasciatus n. sp. and U. heterospinus n. sp. , with data for the latter split into two morphs with seven and eight dorsal-fin spines based on all examined adult specimens.
U. caudofaciatus n. sp. U. heterospinus n. sp.
HT Paratypes HT Paratypes 7 dorsal-fin spines 8 dorsal-fin spines
Morphometric characters Min Mean Max n Min Mean Max n Min Mean Max n Min Mean Max n
Standard length (SL) 116 53 84.7 124 34 91 56 101.4 152 29 66 89.4 122 48 65 90.5 152 53
in % SL
Body depth at first dorsal-fin origin 26 24 26.3 28 34 24 22 23.8 26 29 23 24.1 26 47 22 24.0 26 52
Body depth at anal-fin origin 21 19 21.6 24 34 20 18 20.9 22 29 19 20.7 22 48 18 20.6 22 52
Half body depth at first dorsal-fin origin 21 19 21.3 24 28 20 18 19.9 22 28 18 20.1 22 43 18 19.8 22 43
Half body depth at anal-fin origin 16 15 16.2 18 27 16 15 16.3 17 25 15 16.1 17 40 15 16.0 17 42
Caudal-peduncle depth 10 9.7 10.4 11 34 10 9.8 10.3 11 29 9.2 10.3 11 48 9.2 10.2 11 53
Caudal-peduncle width 3.4 2.6 3.7 4.3 32 4.8 3.6 4.2 5.0 29 3.5 4.2 5.0 47 3.6 4.2 4.8 52
Maximum head depth 22 20 22.2 24 32 21 19 20.9 22 29 19 20.8 23 48 19 20.6 23 53
Head depth through eye 18 16 17.9 20 32 17 15 16.2 17 29 15 16.2 18 48 15 16.0 17 53
Suborbital depth 11 9.6 10.7 13 32 9.9 8.8 9.8 11 29 8.5 9.6 11 48 8.3 9.7 11 53
Interorbital length 8.0 7.5 8.1 8.8 32 8.2 7.3 8.0 9.2 29 7.2 8.1 9.2 48 7.3 8.0 9.0 51
Head length 29 28 29.6 31 32 29 27 28.5 30 29 27 28.7 31 48 27 28.7 31 53
Snout length 12 9.9 11.4 12 32 11 9.9 11.0 12 29 9.9 11.1 13 48 9.7 11.1 12 53
Postorbital length 11 10 11.6 13 32 11 10 11.1 12 29 10 11.2 13 48 10 11.3 13 53
Orbit length 6.8 6.9 7.6 8.5 32 7.2 5.9 7.2 8.2 29 6.3 7.4 8.2 48 5.9 7.3 8.3 53
Orbit depth 5.9 6.0 6.7 7.9 32 6.7 4.9 6.3 7.1 29 5.5 6.5 7.4 48 4.9 6.5 7.3 53
Upper-jaw length 11 9.9 11.5 13 32 10 9.6 10.7 11 29 9.4 10.9 12 48 9.7 11.0 13 53
Lower-jaw length 11 8.6 10.7 12 32 9.5 9.0 10.0 11 29 8.9 10.3 12 48 9.0 10.3 12 53
Snout width 9.1 8.0 8.9 9.7 27 9.2 8.1 9.2 11 28 7.3 8.7 11 46 7.1 8.5 10 52
Barbel length 20 18 20.4 23 34 18 17 18.5 20 29 16 17.9 20 48 16 17.8 20 53
Maximum barbel width 1.0 0.8 0.9 1.1 32 0.8 0.8 1.0 1.2 29 0.8 1.0 1.2 48 0.8 0.9 1.1 53
First pre-dorsal length 36 34 37.2 41 32 37 35 36.0 38 29 34 36.5 38 48 35 36.2 39 53
Second pre-dorsal length 64 61 64.1 66 32 63 61 63.1 65 29 60 63.3 66 48 61 63.3 67 52
Interdorsal distance 15 12 14.2 17 32 15 12 14.6 16 29 13 14.9 17 48 13 14.9 17 52
Caudal-peduncle length 23 21 23.1 25 32 25 22 23.5 25 29 22 23.8 26 48 22 24.1 26 52
Pre-anal length 65 60 64.9 68 32 64 62 63.7 68 29 61 64.1 68 48 61 63.7 68 52
Pre-pelvic length 34 30 33.9 37 32 31 30 31.7 35 29 30 32.0 34 48 30 31.6 35 52
......continued on the next page TABLE 8. (Continued)
U. caudofaciatus n. sp. U. heterospinus n. sp.
HT Paratypes HT Paratypes 7 dorsal-fin spines 8 dorsal-fin spines
Min Mean Max n Min Mean Max n Min Mean Max n Min Mean Max n
Pre-pectoral length 32 29 31.9 35 32 30 29 29.6 32 29 28 30.1 33 48 28 30.0 33 52
Second dorsal-fin depth 21 20 22.5 24 32 21 19 21.4 23 29 19 21.4 23 47 19 21.2 23 52
Pelvic-fin depth 25 24 26.1 28 32 24 22 24.0 26 29 22 24.1 26 47 22 23.9 26 52
Pectoral-fin depth 17 15 17.5 19 32 17 15 16.8 18 29 15 16.6 18 47 15 16.6 18 52
Length of first dorsal-fin base 15 14 15.8 18 32 15 13 15.3 17 29 14 15.3 17 48 13 15.3 17 51
Length of second dorsal-fin base 14 12 14.1 16 32 13 12 14.1 15 29 12 13.8 15 48 12 13.8 16 52
Caudal-fin length 27 27 30.0 32 32 30 27 29.3 32 23 27 29.1 31 43 27 29.5 32 50
Length of anal-fin base 11 9.7 11.6 13 32 11 10 11.4 13 29 9.7 11.3 13 47 9.9 11.4 13 52
Anal-fin height 17 16 17.5 20 32 16 15 16.4 18 29 15 16.7 19 47 15 16.8 19 51
Pelvic-fin length 21 20 22.7 25 34 20 19 20.5 22 29 19 20.9 23 48 19 20.8 23 52
Pectoral-fin length 20 19 21.5 23 32 19 19 20.0 21 25 19 20.5 22 44 19 20.3 22 53
Pectoral-fin width 4.5 3.6 4.3 4.8 32 4.3 3.8 4.2 4.6 29 3.4 4.1 4.7 48 3.4 4.1 4.7 53
First dorsal-fin height 20 20 21.6 23 30 21 18 20.0 22 27 19 20.5 22 45 18 20.3 23 53
Second dorsal-fin height 18 18 19.4 21 34 16 16 17.2 19 27 16 17.8 20 45 16 17.7 20 52
Meristic characters
Dorsal-fin spines 8 8 8.0 8 34 7 7 7.5 8 29 7 7.0 7 48 8 8.0 8 53
Pectoral-fin rays 14 12 13.4 14 34 14 13 13.8 15 29 13 13.8 14 48 13 13.9 15 53
Rudimentary gill rakers on upper limb 3 3 3.2 5 34 3 2 2.8 4 29 2 3.1 4 48 2 2.9 4 53
Developed gill rakers on upper limb 3 2 2.9 4 34 3 2 3.0 4 29 2 2.6 3 48 2 2.7 4 53
Developed gill rakers on lower limb 11 11 12.7 15 34 13 11 12.3 14 29 11 11.9 13 48 11 12.2 14 53
Rudimentary gill rakers on lower limb 6 3 4.9 6 34 3 3 4.4 6 29 3 4.4 6 48 3 4.5 6 53
Total gill rakers on upper limb 6 6 6.1 7 34 6 5 5.8 6 29 5 5.7 6 48 4 5.6 6 53
Total gill rakers on lower limb 17 16 17.6 19 34 16 16 16.8 18 29 15 16.3 17 48 15 16.7 18 53
Total gill rakers 23 22 23.6 25 34 22 21 22.6 24 29 21 22.0 23 48 21 22.3 24 53
Scales along lateral line 29 28 29.0 30 21 29 28 29.2 30 25 28 29.1 30 32 28 29.2 30 34
Colour patterns
Bars on upper caudal-fin lobe 4 0 2.6 6 34 3 0 4.2 6 29 0 3.4 6 48 0 3.6 6 53
Bars on lower caudal-fin lobe 5 0 4.3 6 34 5 1 4.8 6 29 0 4.5 6 48 0 4.3 6 53
Bars on both caudal-fin lobes 9 0 6.9 11 34 8 1 9.0 12 29 0 7.9 12 48 0 7.9 12 53
Pigmentation degree 4 1 3.0 4 33 4 1 3.8 4 29 0 2.9 4 48 0 2.9 4 53
TABLE 9. Morphometric, meristic and preserved colour characters in all examined adults of four margarethae -group species.
U. caudofasciatus n. sp. U. heterospinus n. sp. U. mouthami U. randalli
Morphometric characters Min Mean Max n Min Mean Max n Min Mean Max n Min Mean Max n
Standard length (SL) 67 88.8 124 42 65 90.0 152 101 73 84.1 94 4 66 84.3 106 8
in % SL
Body depth at first dorsal-fin origin 24 26.2 28 42 22 24.1 26 99 21 22.7 24 4 23 23.5 24 8
Body depth at anal-fin origin 19 21.6 24 42 18 20.7 22 100 19 19.9 21 4 19 20.2 22 8
Half body depth at first dorsal-fin origin 19 21.3 24 38 18 19.9 22 86 19 19.7 20 2 19 19.6 20 8
Half body depth at anal-fin origin 15 16.3 18 37 15 16.0 17 82 15 15.3 15 2 15 15.7 17 8
Caudal-peduncle depth 9.9 10.5 11 42 9.2 10.2 11 101 9.1 9.6 10 4 9.9 10.3 11 8
Caudal-peduncle width 2.6 3.7 4.3 42 3.5 4.2 5.0 99 3.2 3.6 3.9 4 2.9 3.4 3.8 8
Maximum head depth 20 22.2 24 42 19 20.7 23 101 20 20.1 21 4 19 20.3 21 8
Head depth through eye 16 17.8 20 42 15 16.1 18 101 17 17.6 18 4 15 15.6 16 8
Suborbital depth 9.2 10.8 13 42 8.3 9.6 11 101 9.0 10.2 11 4 8.0 9.1 10 8
Interorbital length 7.5 8.1 8.8 42 7.2 8.0 9.2 99 7.6 7.8 8.1 4 7.2 8.0 8.8 8
Head length 28 29.5 31 42 27 28.7 31 101 29 29.4 30 4 27 28.8 30 8
Snout length 9.9 11.4 12 42 9.7 11.1 13 101 11 11.4 12 4 11 11.3 12 8
Postorbital length 10 11.7 13 42 10 11.3 13 101 11 11.4 12 4 10 11.1 12 8
Orbit length 6.7 7.6 8.7 42 5.9 7.3 8.3 101 7.8 8.2 8.8 4 6.2 6.9 7.4 8
Orbit depth 5.8 6.7 7.7 42 4.9 6.5 7.4 101 6.6 7.0 7.7 4 5.1 5.9 6.4 8
Upper-jaw length 9.9 11.5 13 42 9.4 10.9 13 101 9.3 10.5 11 4 10 10.9 11 8
Lower-jaw length 8.6 10.7 12 42 8.9 10.3 12 101 8.8 10.3 11 4 9.1 10.0 10 8
Snout width 8.0 8.9 9.7 35 7.1 8.6 11 98 8.9 9.8 11 2 7.8 8.4 9.0 8
Barbel length 18 19.9 23 42 16 17.9 20 101 20 20.8 22 4 16 18.2 20 8
Maximum barbel width 0.8 0.9 1.1 42 0.8 0.9 1.2 101 0.9 1.1 1.4 4 0.6 0.7 0.7 8
First pre-dorsal length 34 37.0 40 42 34 36.3 39 101 37 37.7 39 4 34 36.6 38 8
Second pre-dorsal length 61 64.3 67 42 60 63.3 67 100 62 63.7 65 4 61 64.2 66 8
Interdorsal distance 12 14.5 17 42 13 14.9 17 100 12 13.8 16 4 13 14.4 15 8
Caudal-peduncle length 21 23.1 25 42 22 23.9 26 100 21 22.2 24 4 23 23.6 25 8
Pre-anal length 60 64.7 68 42 61 63.9 68 100 64 64.7 65 4 63 64.6 67 8
Pre-pelvic length 30 33.8 37 42 30 31.8 35 100 28 29.8 33 4 31 32.0 34 8
Pre-pectoral length 29 31.7 34 42 28 30.0 33 100 29 31.7 34 4 29 30.5 32 8
Second dorsal-fin depth 20 22.5 24 42 19 21.3 23 99 19 20.3 21 4 19 20.7 22 8
......continued on the next page TABLE 9. (Continued)
U. caudofasciatus n. sp. U. heterospinus n. sp. U. mouthami U. randalli
Min Mean Max n Min Mean Max n Min Mean Max n Min Mean Max n
Pelvic-fin depth 24 26.0 29 42 22 24.0 26 99 21 23.1 24 4 23 23.4 24 8
Pectoral-fin depth 15 17.5 19 42 15 16.6 18 99 16 16.7 17 4 15 16.0 17 8
Length of first dorsal-fin base 14 15.9 18 42 13 15.3 17 99 11 12.2 13 4 15 15.2 16 8
Length of second dorsal-fin base 12 14.3 16 42 12 13.8 16 100 13 13.5 14 4 12 13.6 15 8
Caudal-fin length 27 29.9 32 42 27 29.3 32 93 27 27.8 29 4 27 28.3 30 8
Length of anal-fin base 9.7 11.6 13 42 10 11.4 13 99 11 12.2 13 4 9.4 10.2 11 8
Anal-fin height 16 17.3 20 42 15 16.8 19 98 17 17.9 19 4 15 16.7 19 8
Pelvic-fin length 20 22.3 24 42 19 20.8 23 100 22 22.3 23 4 20 21.2 22 8
Pectoral-fin length 19 21.3 23 42 19 20.4 22 97 22 22.7 23 4 20 21.1 22 8
Pectoral-fin width 3.6 4.3 4.8 42 3.4 4.1 4.7 101 4.0 4.2 4.3 4 3.8 4.1 4.4 8
First dorsal-fin height 19 21.3 23 39 18 20.4 23 98 20 20.2 21 3 19 20.7 22 8
Second dorsal-fin height 18 19.1 21 38 16 17.8 20 97 17 18.1 19 3 16 17.6 20 8
Meristic characters
Dorsal-fin spines 8 8.0 8 42 7 7.5 8 101 8 8.0 8 4 8 8.0 8 8
Pectoral-fin rays 12 13.5 14 42 13 13.9 15 101 13 13.0 13 4 14 14.0 14 8
Rudimentary gill rakers on upper limb 2 3.1 5 42 2 3.0 4 101 1 2.3 3 4 2 2.9 3 8
Developed gill rakers on upper limb 2 2.9 4 42 2 2.7 4 101 3 4.0 5 4 3 3.3 4 8
Developed gill rakers on lower limb 10 12.4 15 42 11 12.1 14 101 13 13.0 13 4 12 13.5 15 8
Rudimentary gill rakers on lower limb 4 5.1 7 42 3 4.5 6 101 4 4.5 5 4 4 4.6 6 8
Total gill rakers on upper limb 5 6.0 7 42 4 5.7 6 101 6 6.3 7 4 6 6.1 7 8
Total gill rakers on lower limb 16 17.5 19 42 15 16.5 18 101 17 17.5 18 4 17 18.1 19 8
Total gill rakers 22 23.6 25 42 21 22.2 24 101 23 23.8 25 4 23 24.3 25 8
Scales along lateral line 28 29.1 30 29 28 29.2 30 66 29 29.0 29 1 28 29.1 30 8
Colour patterns
Bars on upper caudal-fin lobe 0 2.5 6 42 0 3.5 6 101 0 0.8 2 4 0 0.5 2 8
Bars on lower caudal-fin lobe 0 3.9 6 42 0 4.4 6 101 0 2.3 3 4 0 1.1 4 8
Bars on both caudal-fin lobes 0 6.4 11 42 0 7.9 12 101 0 3.0 5 4 0 1.6 5 8
Pigmentation degree 0 2.7 4 41 0 2.9 4 100 0 1.3 2 4 0 1.8 3 8
TABLE 10. Morphometric, meristic and preserved colour characters in adult U. heterospinus n. sp. , from different areas, with published data from Bandai et al . (2018) for a single specimen from Japan added.
Gulf of Thailand South central Vietnam N Vietnam Indonesia Singapore Philippines Japan
Morphometric characters Min Mean Max n Min Mean Max n Min Mean Max n Min Mean Max n (Bandai et al .)
Standard length (SL) 66 78.6 123 44 70 103.3 152 30 72 96.2 122 20 65 88.4 127 4 78 81 96 96
in % SL
Body depth at first dorsal-fin origin 23 24.4 26 44 22 23.9 26 30 22 23.5 25 20 23 24.0 25 4 - - 24 24
Body depth at anal-fin origin 19 20.7 22 44 19 21.0 22 30 18 20.1 22 20 20 20.7 21 4 22 - 22 21
Half body depth at first dorsal-fin 18 20.2 22 37 18 19.9 22 29 19 19.6 21 16 18 19.8 22 3 - - 20 21
origin Half body depth at anal-fin origin 15 15.9 17 38 15 16.3 17 26 15 15.8 17 14 15 15.3 16 3 - - 17 16
Caudal-peduncle depth 9.4 10.3 11 44 9.8 10.3 11 30 9.2 9.8 11 20 10 10.2 11 4 11 11 11 9.8
Caudal-peduncle width 3.5 4.2 4.7 44 3.6 4.3 5.0 30 3.8 4.3 4.7 20 3.7 3.9 4.1 4 - - 4.1 3.6
Maximum head depth 19 20.9 23 44 19 20.9 22 30 19 19.9 21 20 20 20.8 21 4 21 20 20 20
Head depth through eye 15 16.2 18 44 15 16.3 17 30 15 15.7 17 20 15 15.9 16 4 16 17 15 17
Suborbital depth 8.7 9.7 11.1 44 8.8 9.8 11 30 8.5 9.3 10 20 8.8 9.8 11 4 8.7 8.5 8.3 9.4
Interorbital length 7.5 8.2 9.2 42 7.3 8.0 9.2 30 7.2 7.7 8.2 20 7.3 7.8 8.3 4 8.7 9.0 8.0 7.8
Head length 27 29.0 31 44 27 28.5 30 30 27 28.7 30 20 27 27.8 29 4 28 29 27 30
Snout length 10 11.2 13 44 9.9 11.0 12 30 10 11.0 12 20 10 11.3 12 4 9.9 12 9.7 11
Postorbital length 10 11.4 13 44 10 11.1 12 30 10 11.1 12 20 10 11.5 12 4 11 12 11 12
Orbit length 6.4 7.5 8.3 44 5.9 7.2 8.2 30 6.3 7.3 8.0 20 6.8 7.4 8.0 4 7.3 7.4 7.4 7.3
Orbit depth 5.8 6.7 7.4 44 4.9 6.3 7.0 30 5.5 6.5 7.1 20 6.0 6.5 7.1 4 6.4 5.9 6.6 6.2
Upper-jaw length 10 11.3 13 44 9.6 10.7 11 30 9.4 10.6 11 20 9.8 10.9 12 4 11 12 11 11
Lower-jaw length 9.5 10.6 12 44 9.0 10.0 11 30 8.9 10.0 11 20 9.5 10.2 11 4 10 11 10 10
Snout width 7.1 8.2 11 44 8.1 9.2 11 29 7.9 8.6 9.5 20 7.5 8.1 8.8 4 - - 9.1 8.8
Barbel length 16 17.6 20 44 17 18.4 20 30 16 17.3 20 20 17 18.3 19 4 18 20 17 17
Maximum barbel width 0.8 0.9 1.2 44 0.8 1.0 1.2 30 0.8 0.9 1.1 20 0.8 0.9 1.0 4 0.9 0.9 0.8 1.2
First pre-dorsal length 35 36.7 39 44 35 36.0 38 30 34 36.0 38 20 35 36.3 39 4 36 36 36 37
Second pre-dorsal length 60 63.7 67 44 61 63.2 65 30 61 62.9 64 20 61 62.9 64 4 63 - 64 64
Interdorsal distance 13 15.1 17 44 13 14.6 16 30 13 14.7 16 20 14 14.9 16 4 15 - 16 16
Caudal-peduncle length 22 24.0 26 44 22 23.6 25 30 22 24.1 25 20 24 24.2 25 4 25 - 24 24
Pre-anal length 61 64.3 67 44 62 63.8 68 30 61 63.2 65 20 61 63.2 65 4 68 - 63 64
Pre-pelvic length 30 32.0 35 44 30 31.6 33 30 30 31.5 33 20 31 32.5 33 4 33 - 31 31
......continued on the next page TABLE 10. (Continued)
Gulf of Thailand South central Vietnam N Vietnam Indonesia Singapore Philippines Japan
Min Mean Max n Min Mean Max n Min Mean Max n Min Mean Max n (Bandai et al .)
Pre-pectoral length 29 30.6 33 44 29 29.6 31 30 28 29.4 31 20 28 30.3 32 4 30 - 29 29
Second dorsal-fin depth 20 21.5 23 44 20 21.5 23 30 19 20.6 23 20 20 21.7 23 4 - - 23 22
Pelvic-fin depth 22 24.4 26 44 22 24.1 26 30 22 23.0 25 20 23 23.9 25 4 - - 24 25
Pectoral-fin depth 15 16.7 18 44 15 16.8 18 30 15 16.3 18 20 16 16.5 17 4 - - 16 17
Length of first dorsal-fin base 14 15.2 17 43 13 15.4 17 30 14 15.1 17 20 14 15.2 16 4 16 - 15 15
Length of second dorsal-fin base 12 13.4 15 44 12 14.1 15 30 12 14.2 16 20 14 14.1 15 4 13 - 14 14
Caudal-fin length 27 29.1 30 43 27 29.3 32 24 28 29.7 31 20 29 29.2 30 4 29 - 30 -
Length of anal-fin base 9.7 11.4 13 44 10 11.4 13 30 10 11.2 12 20 10 11.0 12 4 - - 12 11
Anal-fin height 15 17.1 19 43 15 16.3 18 30 15 16.7 18 20 15 17.2 19 4 - - 15 -
Pelvic-fin length 19 21.1 23 43 19 20.4 22 30 19 20.9 22 20 20 20.5 21 4 21 20 21 22
Pectoral-fin length 19 20.6 22 44 19 19.9 21 26 19 20.8 22 20 20 19.8 20 4 20 20 19 21
Pectoral-fin width 3.4 4.0 4.7 44 3.9 4.2 4.6 30 3.7 4.2 4.6 20 3.9 4.3 4.6 4 4.6 4.3 4.0 4.8
First dorsal-fin height 19 20.6 23 43 18 20.0 22 29 19 20.6 23 19 19 19.9 21 4 22 20 21 21
Second dorsal-fin height 16 18.0 20 44 16 17.2 19 28 16 18.1 19 19 16 17.7 19 4 16 - 17 18
Meristic characters
Dorsal-fin spines 7 7.5 8 44 7 7.5 8 30 7 7.6 8 20 7 7.8 8 4 7 8 8 7
Pectoral-fin rays 13 13.8 14 44 13 13.9 15 30 13 14.0 15 20 13 13.8 14 4 13 14 14 13
Rudimentary gill rakers on upper limb 2 3.1 4 44 2 2.8 4 30 2 3.1 4 20 3 3.0 3 4 2 2 2 3
Developed gill rakers on upper limb 2 2.4 3 44 2 3.0 4 30 2 2.7 3 20 3 3.0 3 4 3 3 4 2
Developed gill rakers on lower limb 11 11.8 13 44 11 12.4 14 30 11 12.1 13 20 12 12.3 13 4 12 12 12 13
Rudimentary gill rakers on lower limb 4 4.4 6 44 3 4.4 6 30 3 4.6 6 20 4 4.8 5 4 4 5 5 5
Total gill rakers on upper limb 4 5.5 6 44 5 5.8 6 30 5 5.8 6 20 6 6.0 6 4 5 5 6 5
Total gill rakers on lower limb 15 16.2 18 44 16 16.8 18 30 15 16.6 18 20 17 17.0 17 4 16 17 17 18
Total gill rakers 21 21.7 24 44 21 22.6 24 30 21 22.4 23 20 23 23.0 23 4 21 22 23 23
Scales along lateral line 28 29.0 30 25 28 29.2 30 26 29 29.5 30 11 29 29.3 30 3 0 0 28 -
Colour patterns
Bars on upper caudal-fin lobe 0 2.7 6 44 0 4.1 6 30 1 4.2 6 20 3 4.5 6 4 0 0 6 -
Bars on lower caudal-fin lobe 3 4.1 6 44 1 4.8 6 30 3 4.7 6 20 5 5.5 6 4 0 0 3 -
Bars on both caudal-fin lobes 4 6.9 11 44 1 8.9 12 30 5 8.9 11 20 8 10.0 12 4 0 0 9 -
Pigmentation degree 2 2.4 3 43 1 3.8 4 30 1 3.1 4 20 2 3.0 4 4 0 0 1 -
Etymology. Named after the type locality which is situated close to Gubal Island and in the Strait of Gubal at the southern entrance to the Gulf of Suez, Northern Red Sea . The name “ gubal ” is used as a noun in apposition. Distribution, habitat and size. Single locality off Gubal Island, S Gulf of Suez, Northern Red Sea ; fish caught by trawling at 70 m depth, most probably on sandy or muddy bottom; size of the specimen is 8.7 cm SL. Remarks. Fresh colour of Upeneus gubal n. sp. is yet unknown, but the available morphological evidence strongly suggests its inclusion in the margarethae group (see also the Interspecific comparisons section further below). The single type specimen was collected together with a specimen of Upeneus moluccensis (Bleeker, 1855) (registered as MNHN 2011-0093). The latter did not show any remains of dark pigmentation e.g. of first dorsal-fin tip and oblique bars on upper caudal-fin lobe as indicated to be diagnostic for preserved conspecifics ( Uiblein & Heemstra 2010 ). The single known locality of U. gubal n. sp. in the S Gulf of Suez is about 1100 km north of the northernmost currently known occurrence of U. margarethae off Ibn Abbas Island, S coast of Sudan .